Paryphoconus Enderlein

Borkent, Art, 2014, The Pupae of the Biting Midges of the World (Diptera: Ceratopogonidae), With a Generic Key and Analysis of the Phylogenetic Relationships Between Genera, Zootaxa 3879 (1), pp. 1-327 : 112-113

Paryphoconus Enderlein

( Figs. 12H , 22L , 28I , 31J , 41F , 46T , 54C , 71C , 78L )

DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 8 with D-3-VIII ( Fig. 12H ); also, the female is the only Ceratopogonidae with wings abutting ventromedially ( Figs. 12H , 41F ).

DESCRIPTION: Habitus (female) as in Fig. 12H . Total length = 4.50–6.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B , 33B ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C , 79H ). Head: Dorsal apotome ( Fig. 22L ), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28I ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna ( Fig. 41F ) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), not narrowed sharply posteriorly; sensilla: dorsal apotomals ( Fig. 22L )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 28I )—2 thick setae; oculars ( Fig. 28I )—1 seta, 1 campaniform sensillum? or possibly merely an indentation? Thorax: Prothoracic extension ( Fig. 28I ) wide, well-developed but narrow dorsolaterally, extending from palpus to antenna; mesonotum yes tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 54C ); respiratory organ ( Fig. 46T ) length/width = 3.06–5.06, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single curved row, outer surface with some wrinkles, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ short, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, plates to half length; wing ( Fig. 41F ) without apical tubercle or angle, with males wings separated medially by fore-, midlegs, female wings abutting medially; halter apex and hind leg (as in Fig. 33J ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41F ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33K ); male with apex of foreleg moderately anterior to apex of midleg (as in Fig. 41E ), female with apices of fore-, mid leg not externally visible, dorsal to fused wings; apex of hind leg abutting apex of midleg laterally (dorsal to wing in female); sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (as in Figs. 31L–M ); anterolaterals—1 elongate seta; dorsal setae ( Fig. 31J )—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3- T campaniform sensillum; D-1-T, D-2-T separate or on single tubercle, D-3-T lateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 54C )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe and 2 anterolateral spots or with bare patches in 3 medial and anterolateral areas, sternites 3–7 with medial stripe, anterolateral spot; sternite 8 with dark posteromedial apodeme, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 78L ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 54C ) with 7 setae, 1 campaniform sensillum, including 3 lateral sensilla, D- 2-I, D-3-I closely approximated, D-7-I absent; segment 4 ( Fig. 71C )—D-2-IV, D-3-IV elongate setae on elongate tubercles; D-5-IV, D-8-IV, D-9-IV elongate setae; D-5-IV on single elongate tubercle, D-8-IV, D-9-IV on separate but closely approximated elongate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-8-IV, D-9-IV; D-7-IV near D-3-IV; L-1-IV elongate seta on elongate tubercle, just anterior of base of tubercle with L-3-IV; L-2-IV, L-3-IV, L-4-IV elongate setae on pointed tubercles, V-5-IV, V-6- IV, V-7-IV elongate setae, on short or moderately elongate tubercles, V-6-IV, V-7-IV somewhat closely approximated; segment 8 with D-3-VIII, with or without L-1-VIII; segment 9 ( Fig. 78L )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Paryphoconus is known from 41 species in the Nearctic (one species) and Neotropical Regions ( Borkent 2014 ). Pupae have been collected among vegetation along creeks or on the sandy bottom of a shallow stream.

TAXONOMIC DISCUSSION: The pupae of four species of Paryphoconus are known ( Tables 2–3 ). A female pupa of a Mallochohelea in the ZSMC was labeled " Paryphoconus lanei " but on a modern label and with recent handwriting. The specimen was among material studied by Mayer but does not correspond to his description of Paryphoconus flavidus (as lanei ) ( Mayer 1959b). The Paryphoconus pupae studied by Mayer (1959b) were not located.

MATERIAL EXAMINED: P. grandis : 2 pupal exuviae, Ruta Nacional 12, Corrientes, Argentina, 18-IX- 2010 ( MLPA). P. oliveirai : 1 pupal exuviae, Parque das Garcas, Amazonas Igarape, Brazil, 14-X-2005 ( INPA) .

Figures

FIGURE 12. Habitus. A. Pellucidomyia leei, male, in dorsal view (from Debenham 1970). B. Dibezzia prominens, sex unknown, in dorsal view (form Mayer 1934c). C. Macropeza albitarsis, male, in dorsal view (from Szadziewski & Dominiak, 2007, modified). D. Mallochohelea munda, male, in ventral view (from Szadziewski et al. 1997). E. Sphaeromias fasciata, sex unknown, in dorsal view (from Mayer 1934a). F. Bezzia glabra, female, in ventral view (from Thomsen 1937). G. Palpomyia sp., male, in dorsal view (from Grogan & Wirth 1979). H. Paryphoconus oliveirai, female, in ventral view (modified from Ronderos et al. 2007).

FIGURE 13. Anterior portion of heads and thoraces, in anterior or anterodorsal view. A. Leptoconops kerteszi, ecdysial sutures of head enhanced to show position. B. Culicoides denticulatus. C. Ceratopogon nr. abstrusus. D. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). E. Alluaudomyia bella, dorsal apotome absent. F. Clinohelea curriei. G. Pellucidomyia leei. H. Mallochohelea caudellii. I. Bezzia sp. (from Bolean Lake, BC, Canada).

FIGURE 16. Anterior portion of heads. A. Parabezzia huberti, in ventral view (distorted slide specimen). B. Parabezzia sp. (5 km. E. Danby, Vermont, USA), ventrolateral view. C. Clinohelea curriei, in ventral view. D. Heteromyia clavata, in ventral view, dorsal apotome not present. E. Heteromyia wokei, in ventral, slightly lateral view. F. Pellucidomyia leei, in ventral view, dorsal apotome not present.

FIGURE 17. Anterior portion of heads. A. Jenkinshelea sp., in ventral view (from Ottawa, ON, Canada). B. Mallochohelea caudellii., in ventral view. C. Sphaeromias longipennis, in ventral view. D. Bezzia sp., in ventral view (6 km E Salmon Arm, BC, Canada). E. Palpomyia sp. (Black Lake, Quebec, Canada), in ventral view. F. Phaenobezzia sp., in ventral view (from Skulsuza, South Africa).

FIGURE 22. Dorsal apotomes, anterior view. A. Sphaeromias longipennis (from Wirth & Grogan 1979). B. Amerohelea sordidipes (from Lane et al. 1955). C. Bezzia mollis (from Mayer 1934c). D. Bezzia roldani (from Spinelli & Wirth 1989). E. Bezzia brevicornis (from Spinelli 1983a). F. Clastrieromyia dycei (from Spinelli & Grogan 1986). G. Pachyhelea pachymera (from Spinelli 1983a). H. Palpomyia lineata (from Grogan & Wirth 1979). I. Palpomyia belkini (from Grogan & Wirth 1979, modified). J. Palpomyia flaviceps (from Grogan & Wirth 1979). K. Phaenobezzia opaca (from Wirth & Grogan 1982). L. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 28. Posterior portion of heads, in ventral view. A. Leehelea hispida. B. Sphaeromias longipennis. C. Xenohelea galatea. D. Bezzia nobilis. E. Clastrieromyia dycei. F. Pachyhelea pachymera. G. Palpomyia subasper. H. Phaenobezzia opaca. I. Paryphoconus grandis.

FIGURE 31. Dorsal setae from left side of thoraces; anteromedial setae. A. Homohelea delanoe. B. Leehelea hispida. C. Sphaeromias longipennis. D. Xenohelea galatea. E. Bezzia nobilis. F. Clastrieromyia dycei. G. Pachyhelea pachymera. H. Palpomyia jonesi. I. Phaenobezzia mashonensis. J. Paryphoconus oliveirai. K. Echinohelea lanei, partial lateral view (distorted slide specimen). L. Palpomyia (JAD685), dorsal view. M. Palpomyia (JAD685), dorsolateral view. N. Probezzia seminigra, dorsal view of left side.

FIGURE 33. Cephalothoraces and abdominal segments 1–3, in lateral view. A. Parabezzia sp. (5 km. E. Danby, Vermont, USA). B. Clinohelea curriei. C. Pellucidomyia leei. D. Hebetula tonnoiri. E. Jenkinshelea sp. (from Ottawa, ON, Canada). F. Mallochohelea caudellii. G. Probezzia seminigra. H. Nilobezzia sp. (from Kruger NP, South Africa). I. Sphaeromias longipennis. J. Bezzia sp. (6 km E. Salmon Arm, BC, Canada) K. Palpomyia sp. (Black Lake, Quebec, Canada). L. Phaenobezzia sp. (from Skulsuza, South Africa).

FIGURE 41. Legs, wings and apices of the antennae, in ventral view. A. Bezzia nobilis. B. Clastrieromyia dycei (reconstructed from distorted specimen). C. Pachyhelea pachymera. D. Palpomyia occidentalis. E. Phaenobezzia opaca. F. Paryphoconus oliveirai (reconstructed from distorted specimen).

FIGURE 46. Respiratory organs, most in dorsal view, Figs. 46G. in lateral view, pattern on tracheal tubes shown only in some. A. Homohelea albitudinis (from de Meillon & Wirth 1981). B. Homohelea delanoe (from de Meillon & Wirth 1981). C. Leehelea wasselli (from Elson-Harris 1987). D. Leehelea hispida (from Elson-Harris 1987). E. Sphaeromias longipennis (from Wirth & Grogan 1979). F. Xenohelea galatea. G. Amerohelea sordidipes (from Lane et al. 1955). H. Bezzia glabra (from Wirth 1983b). I. Bezzia obelisca (from Wirth 1983a). J. Bezzia dorsasetula (from Wirth 1983a). K. Bezzia turkmenica (from Glukhova 1979b). L. Clastrieromyia dycei (from Spinelli & Grogan 1986). M. Pachyhelea pachymera (from Spinelli 1983a). N. Palpomyia lineata (from Grogan & Wirth 1979). O. Palpomyia jonesi (from Grogan & Wirth 1979). P. Palpomyia belkini (from Grogan & Wirth 1979). Q. Palpomyia rufa (from Grogan & Wirth 1979). R. Palpomyia flaviceps (from Grogan & Wirth 1979). S. Phaenobezzia opaca (from Wirth & Grogan 1982). T. Paryphoconus oliveirai (from Ronderos et al. 2007).

FIGURE 54. Metathoraces and tergites 1, in dorsal view. A. Palpomyia jonesi. B. Phaenobezzia opaca. C. Paryphoconus oliveirai.

FIGURE 71. Abdominal segments four, in dorsal and ventral view (shagreen not shown). A. Palpomyia occidentalis. B. Phaenobezzia opaca. C. Paryphoconus oliveirai.

FIGURE 78. Abdominal segments 9. A. Palpomyia lineata, male, in dorsal and ventral view (from Grogan & Wirth 1979). B. Palpomyia lineata, female, in dorsal and ventral view (from Grogan & Wirth 1979). C. Palpomyia jonesi, male, in dorsal and ventral view (from Grogan & Wirth 1979. D. Palpomyia jonesi, female, in dorsal and ventral view (from Grogan & Wirth 1979). E. Palpomyia belkini, male, in dorsal and ventral view (from Grogan & Wirth 1979). F. Palpomyia belkini, female, in dorsal and ventral view (from Grogan & Wirth 1979). G. Palpomyia rufa, male, in dorsal and ventral view (from Grogan & Wirth 1979). H. Palpomyia flaviceps, female, in dorsal and ventral view (from Grogan & Wirth 1979). I. Phaenobezzia vacunae, male, in ventral view (from Haeselbarth 1965). J. Phaenobezzia mashonensis, male, in ventral view (from Haeselbarth 1965). K. Phaenobezzia opaca, female, in dorsal and ventral view (from Wirth & Grogan 1982). L. Paryphoconus oliveirai, female, in dorsal view (from Ronderos et al. 2007).

FIGURE 79. Evolution of ecdysial sutures and splits in pupal exuviae of Ceratopogonidae, head and anterior portion of thorax in ventral view. Numbers refer to character states of characters 2 and 8 in text. Red indicates ecdysial splits. A. Leptoconopinae. B. Forcipomyiinae. C. Dasyheleinae. D. Most Ceratopogonini. E. Most Ceratopogonini, Clinohelea. F. Allohelea, Atyphohelea, Parabezzia. G. Heteromyia, Pellucidomyia, Hebetula. H. most Sphaeromiini s. lat., most Palpomyiini, Paryphoconus; some Bezzia are nearly completed fused (as in Fig. 79I), with only the dorsal apotome separate, along with an ecdysial split along the base of the antenna and the dorsal thoracic split. I. Jenkinshelea, Macropeza, Mallochohelea, Probezzia, Neobezzia.